Revision of the Lispe Longicollis-Group (Diptera, Muscidae)

A peer-reviewed open-access journal ZooKeys 235: 23–39 (2012)Revision of the Lispe longicollis-group (Diptera, Muscidae) 23 doi: 10.3897/zookeys.235.3306 Research article www.zookeys.org Launched to accelerate biodiversity research

Revision of the Lispe longicollis-group (Diptera, Muscidae)

Nikita E. Vikhrev1,†

1 Zoological Museum of Moscow University, Bolshaya Nikitskaya 6, Moscow, 125009, Russia (ZMUM)

† urn:lsid:zoobank.org:author:17508957-C0E9-41C9-B22C-D9AF5A691F53

Corresponding author: Nikita E. Vikhrev ([email protected])

Academic editor: M. Hauser | Received 29 April 2012 | Accepted 26 October 2012 | Published 31 October 2012

urn:lsid:zoobank.org:pub:E89C1914-6AC3-48DE-BF7C-F83778E1EF97

Citation: Vikhrev NE (2012) Revision of the Lispe longicollis-group (Diptera, Muscidae). ZooKeys 235: 23–39. doi: 10.3897/zookeys.235.3306

Abstract TheLispe longicollis species-group is revised. Lispe ethiopica sp. n. is described. The following 3 new syno- nyms are established: Lispe assimilis Wiedemann, 1824 (syn: cyrtoneurina Stein, 1900 and modesta Stein, 1913); Lispe manicata Wiedemann, 1830 (syn: forficata Kurahashi & Shinonaga, 2009). Female of Lispe microptera Seguy, 1937 is described for the first time. Identification key for known Eurasian and African species is given.

Keywords Lispe longicollis species-group, Muscidae, Diptera, key, new species, new synonym

Introduction

The Lispe longicollis species-group was proposed by Hennig (1960) based on the char- acteristic shape of a vein M which is distinctly curved forward at apex. The species of this group also share these additional characters: t3 with submedian av, ad and pd setae; abdomen with large, more or less fused trapezoid spots; frontal triangle narrow; dc 2+4, usually only 2 posterior pairs are strong (in L. glabra Wiedemann and L. manicata Wiedemann dc setae should be described as 0+2, but careful examination shows that 4 anterior pairs of minute dc setulae are normally present but broken in most specimens). Hennig divided the group into two subgroups.

Copyright Nikita E. Vikhrev. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 24 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Subgroup 1 included the species with ventral seta on t2 and consisted of L. longicollis Meigen, 1826 (S Palaearctic) and L. cilitarsis Loew, 1856 (North Africa and Near East). In this paper another three species are added to subgroup 1: L. microptera Seguy, 1937 known from Pakistan and India and two Afrotropical species, L. barbipes Stein, 1908 and the here described L. ethiopica sp. n. The main subgroup character is the presence of fine hairs on the meron above the hind coxa; other characters: t2 with ventral seta (except for L. microptera); male hind basitarsus curved and bears long ventral hairs (except for L. longicollis); halves of cercal plate of a subquadrate shape and strongly conjoined with each other (less so in L. longicollis); the flies inhabit banks of salted to fresh water. Hennig’s subgroup 2 included widespread L. assimilis Wiedemann, 1824 and Afri- can L. nuba Wiedemann, 1830 which lack the ventral seta on t2. In this paper another three Oriental species are added to the subgroup 2: L. glabra Wiedemann, 1824, L. manicata Wiedemann, 1830 and L. pacifica Shinonaga & Pont, 1992. Subgroup 2 is characterized as follows: meron bare; t2 without ventral seta; male hind basitarsus unmodified; halves of cercal plate of subtriangular shape and less conjoined with each other; the flies inhabit banks of fresh water only. This revision considers Palaearctic, Afrotropical and Oriental species of the Lispe longicollis group. The group is absent in the Nearctic region, there are two Australian species not seen by the author, namely Lispe weschei Malloch, 1922 and Lispe xeno- chaeta Malloch, 1923, which also belong to the same group.

Material and methods

The majority of the specimens studied are deposited in the Zoological Museum of Lo- monosov Moscow State University, Moscow, Russia, in this case not indicated in text. Other collections are abbreviated as follows:

BMNH Natural History Museum, London, UK; TAUI Tel-Aviv University, Israel; ZIN Zoological Institute of the Russian Academy of Sciences, St Petersburg, Russia; ZMHU Museum für Naturkunde, Humboldt-Universität zu Berlin, Berlin, Germany.

The names of the collectors are abbreviated as follows: KT – Konstantin Tomko- vich, NV – Nikita Vikhrev. The following abbreviations for morphological structures are used: f1, t1, f2, t2, f3, t3 = fore-, mid-, hind- femur or tibia; ac = acrostichal setae; dc = dorsocentral setae; a, p, d, v = anterior, posterior, dorsal, ventral seta(e); prst – presutural, post - postsutural. The abbreviation for the tarsi as tar followed by a pair of digits separated by a hy- phen was proposed by Vikhrev (2011): the first digit (1 to 3) gives the leg number and the second digit (1 to 5) the number of the tarsal segment. For example, tar1-4 = 4th segment of fore tarsus; tar3-1 = hind basitarsus. Revision of the Lispe longicollis-group (Diptera, Muscidae) 25

Geographical coordinates are given in the Decimal Degrees format. Synonymies are listed only for the species to which the new synonymies are considered, for full lists of synonymies see the regional Diptera Catalogues: Pont 1977, 1980 and 1986.

Identification key to Eurasian and African species of Lispe longicollis-group 1 Meron bare; t2 with only 1 pd seta, without ventral seta. ♂: hind basitarsus not modified as below. Subgroup 2...... 2 – Meron setulose above hind coxa. t2 with ventral seta (exception L. microptera). ♂: hind basi-tarsus modified: curved and with long ventral hairs (exception L. longicollis). Subgroup 1...... 6 2 Disc of scutum densely brownish-grey dusted; dc 2+4, two last prescutellar pairs strong, others at least clearly distinct; presutural intraalar seta present; medium size species, body length 6–7.5mm...... 3 – Disc of scutum mostly subshining, with a pair of thinly dusted submedian vittae only; dc 0+2, only last pair strong; presutural intraalar setae absent; large species, body length 8–9.5mm...... 5 3 ♂: f2 with setae on av to pv surfaces long and dense, the longest setae about twice as long as femur width. ♀: either f2 in basal part with erect, rather dense setulae on av to pv surfaces, these setae at base almost equal to femoral width (L. pacifica) or f1 ventrally with 2–3 rows of fine setulae (L. nuba)...... 4 – ♂: f2 without av-setae and with only short pv-setulae which even in basal part about half as long as femur width. ♀: f2 with only short and sparse setulae; f1 bare on ventral surface apart from usual row of av setae. Africa, Palearctic and Oriental regions, Australia...... assimilis Wiedemann 4 ♂: f1 ventrally with a dense brush of setulae placed in about 5 rows in basal half of femur and in 1–2 rows in apical half, the usual av row of setae on f1 reduced to 1(2) setae at apex. f2 with ventral setae long in basal 1/3 (2 times as long as femur width), ventral setae in median 1/3 much shorter, only as long as femur width. ♀: f1 ventrally with 2-3 rows of fine setulae. Africa...... nuba Wiedemann – ♂: f1 ventrally without setulae; a complete av row on f1 present, though con- sists of fine setae. f2 with ventral setae of equal length in basal 2/3 of femur, about 1.5–2 times as long as femur width. ♀: f1 bare on ventral surface apart from usual row of av setae. East Asia...... pacifica Shinonaga & Pont 5 Palpi darkened at apex. Parafacials with usual sparse fine hairs. f3 with submedian av seta long (equal to femur width) and placed beyond middle Ab- dominal dusted median vitta complete, although vague on anterior parts of tergites. Hind basitarsus without v seta at base. ♂: mid legs and wing venation modified as in Figs 15 and 16; cercal plate as shown in Fig. 11. Oriental region...... glabra Wiedemann 26 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

– Palpi entirely yellow. Parafacials entirely bare. f3 with submedian av seta short (half as long as femur width) and placed before middle. Abdomen with dusted median stripes conspicuous only on posterior half of tergites. Hind basitarsus with a strong “Anthomyiidae-like” v seta at base. ♂: mid tarsi modified as in Fig. 14; wing venation similar to females (Fig. 13); cercal plate as in Fig. 12. South of the Oriental region...... manicata Wiedemann 6 t2 without av seta (L. microptera – 1 pd; ♂ L. barbipes – 1 v-pv; ♀ L. barbipes – 1 pd, 1 v-pv)...... 7 – t2 with 1 p(pd) and 1 av setae...... 8 7 t2 with 1 p(pd) seta. ♂ (Fig. 3): f3 with 4–5 fine longpv in basal half and 1(2) av in basal 1/3; tar3-1 slightly laterally compressed and outward curved, with waved ventral setulae more dense at base and at apex; tar3-2 with waved v setulae; cercal plate as in Fig. 8. ♀: f3 on av surface usually with a short av seta before middle (in some specimens this seta absent). Pakistan, India...... microptera Seguy – t2 with 1 v(pvv) seta. ♂ (Fig. 1): f2 with 2-3 strong, straight and long ventral spines; f3 in basal 1/3 with 1-2 av and 1 long fine pv; t3 at apical 1/3 with long waved setae on ad to av surface; tar3-1 elongated, downward curved; with waved v setulae. ♀: f3 without av seta; t2 with 1 v-pv and 1 pd approximated setae. Africa...... barbipes Stein 8 f3 with a strong submedian av seta. Meron with 6-8 setulae above hind coxa and 2-4 below posterior spiracle and usually with 1 setula present on katepimeron. ♂: hind tarsus simple; cercal plate as in Fig. 3. South Palaearc- tic from Central Europe to Transbaikalia...... longicollis Meigen – f3 without submedian av. Meron only with 4-5 setulae above hind coxa. ♂: hind tarsus modified...... 9 9 Palpi black. ♂ (Fig. 4): mid tarsus simple; tar3-1 dorso-ventraly flattened, distinctly wider than width of t3; cercal plate - Fig. 5, sternite 5 – Fig. 6. Ethiopia...... ethiopica sp. n. – Palpi yellow. ♂ (Fig. 2): mid tarsus with a row of curled setulae on p surface; tar3-1 not widened, less wide than width of t3; cercal plate as in Fig. 7. Near East, N Africa...... cilitarsis Loew

Taxonomy Lispe assimilis Wiedemann, 1824 http://species-id.net/wiki/Lispe_assimilis Figs 10, 19

Lispe quadrilineata Macquart, 1835. Lispe incerta Malloch, 1925. Lispe inexpectata Canzoneri & Meneghini, 1966. Revision of the Lispe longicollis-group (Diptera, Muscidae) 27

Lispe cyrtoneurina Stein, 1900: 393 syn. nov. Type locality: Papua New Guinea, Dilo. Lispe modesta Stein, 1913: 557 syn. nov. Type locality: Abyssinia, Dambelsee [= Ethio- pia, Ziway Lake].

Material examined. Syntype Lispe modesta Stein, 1913 ♂, (ZMHU). [Ethiopia] Ab- yssinia, Lac. Dembel [Ziway Lake], I.1912, Kovacs. Australia,: Qld., Townsville, 19.29°S, 146.80°E, 17.IV.2012, G.Cocks, 1♀. Ethiopia: Amhara, Tana Lake env., 1800m asl, 11.54°N, 37.39°E, 2–4.VIII.2012, NV, 3♂♂, 1♀; Oromia, Ziway Lake, 7.91°N, 38.73°E, 12.III.2012, NV, 1♂, 1♀. India: Goa state, 15.0°N 74.1°E, 3–16.II.2008, KT, 29♂♂♀♀; Rajasthan state, Jaipur, 26.96°N, 75.85°E, 22.II.2011, NV, 7♂♂, 11♀♀; Uttarakhand state, 30.1°N, 78.2°E, 4.IX.2011, NV, 1♀. Israel, Kinneret Lake env., 27.X.2011, NV, 7♂♂, 2♀♀. [Italy], Sicilia, Partinico L., 12.VIII.1978, S.Canzoneri, 1♀, (labeled L. inexpectata) (ZMHU). Myanmar, Shan state, Inle Lake, 30.XI.2009, NV, 6♂♂, 2♀♀. Morocco: Essaouira prov., Essaouira env., 27.III.2009, NV, 1♂, 3♀♀, 1–5.V.2012, NV, 1♂, 2♀♀; Marrakech prov., Marrakech, 21.III.2009, NV, 1♂, Tat-Tan prov, Draa R., 11.V.2012, NV, 1♀. Nigeria, Zungeru (9.81°N 6.16°E), 25.II.1911, J.Macfei, 1♀ (BMNH). Sudan, 08.III.1929, 1♂ with Emden’s identification label L. modesta (BMNH). Thailand: Chiang Mai prov., Sop Poeng env., 17.XI.2009, NV, 1♂; Mae Hong Son prov., Pai env., 19.4°N, 98.4°E, 15–25.XI.2010, NV, 14♂♂. Turkey: Adana prov., Yumurtalik env., IV.2010, NV, 1♂, 1♀; Antalya prov., Manavgat env., IX.2006–9, NV, 16♂♂, 10♀♀; Hatay prov., Samandag env., IV.2010, NV, 3♂♂, 1♀; Mersin prov., Silifke env., IV.2010, NV, 1♂, 1♀; Sakarya prov., Karasu env., V.2009, NV, 1♂; Zonguldak prov., Alapli env., V.2009, NV, 1♂. Distribution. S Palaearctic, Afrotropical and Oriental regions, Australia, Oceania. Synonymies. The taxonomy of L. assimilis was considered by Shinonaga and Pont (1992). In that paper the synonymy of L. quadrilineata, L. incerta and L. inexpectata with L. assimilis was established and the related Oriental species with long ventral hairs on mid femur was described as Lispe pacifica Shinonaga & Pont, 1992, it was shown that L. assimilis in the sense of old authors is L. pacifica, while later authors followed this misinterpretation. Lispe cyrtoneurina Stein, 1900 – syn. nov. of L. assimilis. Stein’s (1900) original description completely fits L. assimilis, the only difference found is 3 (instead of 4) post dc. The male lectotype of L. cyrtoneurina (stored in Genoa, Museo Civico di Storia Naturale di Genova) was reexamined by Adrian Pont. The lectotype is in poor condition, damped and mostly squashed; 4 post dc; everything else fit L. assimilis (Pont, pers. com. and unpublished notes). Lispe modesta Stein, 1913 – syn. nov. of L. assimilis. The very short Stein’s (1913) description fits L. assimilis. Examined by me specimens from Asia and Africa were found similar, the specimens from Ziway Lake in Ethiopia are especially interesting 28 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012) as it is the type locality of L. modesta. In a later paper (Stein, 1918: 175) Stein himself listed L. assimilis from Rangoon (Yangoon, Myanmar) as “Lispe assimilis Wied. var. modesta Stein” and wrote that the male of L. assimilis var. modesta (=L. assimilis in the present interpretation) differs from L. assimilis (=L. pacifica in the present interpretation) only by the absence of long ventral hairs on f2. Lispe pacifica Shinonaga & Pont, 1992. According to the remark cited above Stein in 1918 started to regard L. assimilis and L. pacifica as variations of the same species. In fact, the separation of these species in female sex is sometimes doubtful and males have similar genitalia. Note also that in both species the pollinosity is very variable: dusting on face, parafacialia and parafrontalia from pure white to deep yellow, dusting on parafrontalia and frontal triangle from weak to strong, dusting of scutum from grey to brown, the colour of the tibiae from almost entirely yellow to almost entirely dark. I would like to report that my observations made around Pai (Thailand, Mae Hong Son province) somewhat support the valid taxonomic status of Lispe pacifica Shinonaga & Pont, 1992. Pai town is so far the only locality I know where both species L. assimilis and L. pacifica were found together at the same time and usually at the same pools. A series of 17 males of L. pacifica and 14 males of L. assimilis were collected. All examined males have distinct characters either of one or other species, with no intermediate specimens recorded. Thus in a sympatric condition no trace of crossbreeding between the two species has been found. So, Lispe assimilis Wiedemann, 1824 = Lispe cyrtoneurina Stein, 1900 syn. nov. = Lispe modesta Stein, 1913, syn. nov.

Lispe barbipes Stein, 1908 http://species-id.net/wiki/Lispe_barbipes Fig. 1

Material examined. Syntypes 1♂, 1♀ (ZMHU). S. W. Afrika, Luderitzbucht [Namibia, Luderitz, 26.65°S, 15.16°E], S. Schultze 1♂; S. W. Afrika, Kalahari, Moocane, Wasser- spiegel [Botswana, Mookane, 23.7°S 26.6°E, water level], X.1904, S. Schultze 1♀. As it was reported by Pont and Werner (2006): “there must be some doubt as to whether this is actually a syntype, since the locality [of ♂ syntype] was not mentioned by Stein (1908) and is on the coast of Namibia rather than at the eastern edge of the Kalahari desert in Botswana.” Ethiopia, Afar, Mille env., 530m asl, 11.381°N, 40.731°E, 9.VIII.2012, NV, 1♀. South Africa, [Northern Cape prov.], Olifantshoek [≈27.94°S, 22.74°E], 26.II.1988, D.Simon, 2 ♂♂ (TAUI). [Namibia], South Africa, Van Zylserus [Kunene reg., Van Zyls pass, 17.64°S, 12.71°E, 1000m asl], 12.I.1988, D.Simon, 3 ♂♂ (TAUI and ZMUM). Distribution. Afrotropical: Botswana, Ethiopia, Namibia, South Africa. Redescription. Male. Body size – 7-8 mm. Head with frontal triangle narrow. Parafacial covered with hairs. Antenna black. Arista in basal half with hairs slightly Revision of the Lispe longicollis-group (Diptera, Muscidae) 29

Figure 1–3. ♂♂ L. barbipes Stein (1) L. cilitarsis Loew (2) L. microptera Séguy (3). shorter than antenna width, in apical half bare. Palpus yellow. Scutum and scutellum brownish dusted with indistinct vittae, pleura brownish-grey dusted. dc 2+4 (strong- strong+medium-medium-strong-strong); intraalars 1+2; supraalars 1+2, the posteri- or one weak. Meron with setulae above hind coxa. Wing with vein R4+5 distinctly curved forward. Legs dark, but femora at apex and tibiae in basal half yellow. f1 with a row of 6-7 strong av setae in apical 3/5. t1 with a row of 7-8 short but strong d setae and with submedian p seta. f2 with 2(3) strong, straight and long (2 times as long as femur width) ventral spines; other setae: 1(2) median a seta(e), 2 p at apex. t2 with 1 submedian v seta (which is slightly shifted from true v position onto posterior surface and may be named “pvv seta”). f3 curved; with 1-2 av setae and 1 long but fine pv in basal 1/3, pv preapical present, av preapical absent. t3 with submedian 1 av, 1 ad and 1 pd setae; below middle with a row of 3-4 straight ad; at apical 1/3 with long waved setae on ad to av surface. Hind tarsus modified: tar3-1 elongated, downward curved; with waved ventral setulae, these in apical 1/3 especially long; tar3-2 thickened. Abdo- men grey dusted with large lateral black spots, these on tergites 3 and 4 separated by grey vitta, on tergite 5 fused. Females differs from male as follows: spines on f2 absent; t2 with 2 approximated submedian setae, shorter p-pd and longer v-pv, f3 without av and pv setae (these characters did not mention in Stein’s (1908) original description), t3 without long setae at apex; hind tarsus unmodified.

Lispe cilitarsis Loew, 1856 http://species-id.net/wiki/Lispe_cilitarsis Figs 2, 7

Material examined. Syntype ♂, ZMHU, also seen by Hennig (1960: 426), [Egypt] Assyud [Asyut], Frauenfeld, 1♂. Egypt: Sinai, 21.V.1981, A.Freidberg, 1♂ (TAUI); Cairo, 2♂♂, 1♀ (ZIN); Cai- ro, Port Said, Suez, Luxor, Aswan, 12♂♂, 6♀♀ (ZMHU). 30 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Ethiopia: Amhara, Tana Lake env., 1800m asl, 11.54°N, 37.39°E, 2-4.VIII.2012, NV, 2♂♂; Oromia, Ziway L., 1640m asl, 7.91°N, 38.73°E, 11–13.III.2012, NV, 1♂. Israel: Ma’agan Michael, 28.VII.1964, A.Valdenberg, 19♂♂, 20♀♀ (TAUI); Eilat env., 24.X.2011, NV, 10♂♂, 9♀♀. Morocco: Tan-Tan prov., Draa R., 28.528°N, 10.947°W, 11.V.2012, NV, 1♂, 1♀. Distribution. Egypt, Ethiopia, Israel, Morocco. Also reliably known from Saudi Arabia and Oman (Pont 1991). In Ethiopia L. cilitarsis seems uncommon and re- stricted to northern regions in comparison with resembling L. ethiopica sp. nov., so specimens from Africa should be re-examined and so far I regard other Afrotropical records as doubtful.

Lispe ethiopica Vikhrev, sp. n. urn:lsid:zoobank.org:act:8412D7E2-3B7E-4527-A0F2-6B3DACBEB089 http://species-id.net/wiki/Lispe_ethiopica Figs 4, 5, 6

Holotype: male, Ethiopia, Oromia, Langano Lake, 1590m asl, 7.646°N, 38.706°E, 13-15.III.2012, NV (ZMUM). Paratypes 23♂♂, 24♀♀. Ethiopia: Dire Dawa, Afrika, Diredaua [= Ethiopia, Dire Dawa, 9.60°N, 41.85°E], 28.X.[1945–55], O.Theodor, 1♂(TAUI); Oromia: Mojo bridge, 8.597°N, 39.111°E, 21.IX.2003, A.Freidberg, 1♂ (TAUI); Langano Lake, 1590m asl, 7.646°N, 38.706°E, 13–15.III.2012, NV, 9♂♂, 12♀♀; Ziway Lake, 1640m asl, 7.91°N, 38.73°E, 11–13.III.2012, NV, 11♂♂, 12♀♀; Abijata Lake, 1580m asl, 7.61°N, 38.65°E, 14.III.2012, NV, 1♂. Description. Male, body length 6.5–7.5 mm. Head. Frontal triangle remarkably narrow, brownish in posterior half, yellowish- grey dusted in anterior half. Interfrontalia blackish-brown. Fronto-orbital plate blak- ish-brown in posterior third, yellowish-grey dusted anteriorly; with 3-5 inclinate and 2 proclinate setae and dense hairs in outer row. Parafacial and cheek whitish dusted, occiput grey, parafacial with a row of hairs. Antenna black, postpedicel short, only 2 times longer than pedicel. Arista with hairs half as long as antenna width. Vibrissae medium strong. Palpi blackish. Thorax. Pleura densely grey dusted, scutellum and disc of scutum brown, thinly dusted, with a pair of densely dusted prescutellar ochrous spots; vittae indistinct. Presutural ac in 4 irregular rows; dc 2+4, four anterior pair medium strong, two posterior pairs strong; intraalars 1+2; supraalars 1+2; katepisternals 1+2; anepimeron with 11-13 setulae; meron with 3-5 setulae above hind coxa. Wings hyaline, slightly brownish, vein M distinctly curved forward at apex, calypters white, halter yellow. Legs black with grey dusting, but knees and base of tibiae yellowish. f1 with a row of pd setae and a row of pv setulae; t1 with submedian p seta. f2 with a seta at middle and 2 pd preapicals; t2 with p seta at middle and av seta in apical third; mid tarsus simple. f3 with 1-2 fine v setulae at base, at apex with 1 short av and 1 short pv; t3 Revision of the Lispe longicollis-group (Diptera, Muscidae) 31

Figure 4–6. L. ethiopica sp.n.: male Holotype (4) cercal plate (5) sternite 5 (6). with submedian ad and pd setae and with long fine av at apical third, setulae in the ad row elongated. Hind tarsus modified: tar3-1 dorso-ventrally flattened, distinctly wider than width of t3, on av surface with a dense row of fine curled setulae. Abdomen with dense whitish dusting; tergites 3 to 5 with a pair of large black fused spots each. Cercal plate and sternite 5 as in Figs 5 and 6. Female differs from male as follows: body length 7-8 mm; t3 with av seta strong; hind tarsus simple. Diagnosis. Lispe ethiopica sp.n. is related to L. cilitarsis Loew, 1856 and probably was overlooked due to that resemblance. These two species may be reliably distin- guished in both sexes as recommended in the identification key above. Etymology. Named after the locality of the type series.

Lispe glabra Wiedemann, 1824 http://species-id.net/wiki/Lispe_glabra Figs 11, 15, 16

Material examined. India, Goa state, 15.0°N, 74.1°E, 3–16.II.2008, KT, 3♂, 7♀♀. Myanmar, Shan state, Inle L., 30.XI.2009, NV, 3♀♀. Thailand: Chanthaburi prov., Khao Khitchakut env., 12.82°N, 102.13°E, XI.2009, NV, 3♀♀; Chonburi prov., Pattaya env., XII.2008–9, NV, 40 ♂♂, ♀♀; Mae Hong Son prov., Pai env., 11.XI.2009, NV, 5♀♀; Phuket prov., Nai Thon beach, 20.II.2009. NV 3♂♂, 4♀♀, NV; Phang Nga prov., Thai Mueang env., 18.II.2009, NV, 1♂, 6♀♀; Rayong prov., Ban Phe env., 12.64°N, 101.46°E, NV, 3♀♀. Distribution. Oriental region. 32 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Figure 7–9. Cercal plates: L. cilitarsis Loew (7) L. microptera Séguy (8) and L. longicollis Meigen (9).

Descriptive notes. Body length 8.5–9.5 mm. Wings slightly brownish infuscated. Vein M gradually curved forward from level of crossvein dm-cu, cell r4+5 is almost closed and distance between veins M and R4+5 at wing margin is shorter than crossvein rm. Vein CuA2 not reaching wing margin, extending only to crossvein dm-cu; crossvein dm-cu skewed, it reaches vein M at acute angle of about 45˚. There is a downcurved fold surrounded by long microtrichia along posterior margin of wing between veins M and A2, microtrichia directed outward to the fold. Mid legs: f2 with remark- able row of very dense curled Velcro fastener-like setae in pv position in basal 2/3; t2 in apical 1/4 with a row of long ventral hairs; tar2-1 with a complete row of long curved pv setulae. Male is unmistakable due to modified wings and mid legs. Female differs from female of L. manicata as given in the key.

Lispe longicollis Meigen 1826 http://species-id.net/wiki/Lispe_longicollis Fig. 9

Material examined. [Iran], Sistan [Sistan and Baluchestan prov., ≈27°N, 61°E], 19–21.V.1898, Zarudniy, 2♂♂ (ZMHU). Hungary, Kalocsa, [46.5°N 19.0°E] , Thalhammer, 2♀♀ (ZMHU). Kazakhstan: Atyrau reg., Atyrau env., 47.0N, 51.8E, 21.V.2011, KT, 21♂♂, 17♀♀; Kyzylorda prov., Syr Darya R., KT, 26 ♂♂, ♀♀, W. Kazakhstan reg., Uralsk env., 51.07°N, 51.05°E, 26.VIII.2012, KT, 6♂♂, 7♀♀. Russia: Astrakhan reg., Baskunchak L., 48.19°N, 46.82°E, 2–4.V.2010, KT, 7♂♂, 2♀♀; Gorno-Altay reg., Ust-Koksa env., 50.26°N, 85.61°E, 25.VI.2007, O.Kosterin, 1♀; Kalmykia reg., 47.875°N, 44.601°E, 08.VI.2012, NV, 3♂♂, 1♀; Khakassia reg., Shira env. 54.422°N, 90.147°E, 26.VI.2011, KT, 2♀♀; Kras- nodar reg., Pshada R., 44.39°N, 38.34°E, 6.IX.2009, KT, 6♂♂, 3♀♀; Omsk reg., Omsk, 54.97°N, 73.36°E, 15.VI.2011, O.Kosterin, 1♀; Orenburg reg., Sol- Iletsk env., 51.342°N, 55.013°E, 28.VIII. KT, 7♂♂, 7♀♀; Saratov reg., Saratov env., 51.60°N, 46.35°E, 24.VIII.2012, KT, 1♂, 4♀♀; Stavropol reg.,saltish pond, 45.245°N, 42.665°E, 09.VI.2012, NV, 4♂♂, 2♀♀; Volgograd reg., 50.418°N, Revision of the Lispe longicollis-group (Diptera, Muscidae) 33

Figure 10–12. Cercal plates: L. assimilis (10) L. glabra Wiedemann (11) and L. manicata Wiedemann (12).

42.760°E, 7.VI.2012, NV, 1♂, Sarpa L., 48.35°N, 44.61°E, 7.VI.2012, NV, 3♂♂, 2♀♀; Zabaykalsky reg., Zun-Torey soda Lake, 50.01°N, 115.72°E, 30.VII.2011, A.Medvedev, 1♂. [Slovenia], Illyria, Gorz [Gorica, ≈ 45.9°N 13.7°E], IX.67, Mik, 1♂ (ZMHU). Tajikistan, Khatlon prov.; Jilikul env. (37.5°N, 68.5°E), 16.V.1987, M.Krivosheina, 2♂♂, 2♀♀; Turkey: Antalya prov., Manavgat env., 36.76°N, 31.45°E, X.2006–7, NV, 1♂, 2♀♀; Hatay prov., Samandag env., 36.07°N, 35.96°E, 16.IV.2010, NV, 1♂; Kayseri prov., Subashi env., 38.51°N, 35.19°E , 19.IV.2010, NV, 1♀; Konya prov., Beyshehir Lake, 37.79°N, 31.64°E, 11.IX.2009, NV, 18 ♂♂, ♀♀; Mersin prov., Sil- ifke env., 36.31°N, 34.02°E, 22.IV.2010, NV, 2♂♂, 8♀♀. Turkmenistan: Balkan prov., Atrek R., (37.7°N, 54.8°E), 28.VII.1932, Ushin- sky, 1♀; Mary prov.; Badhyz NR (35.7°N, 61.8°E), V-VI.1991, A.Ozerov, 1♂, 6♀♀. Ukraine, Donetsk reg., Volnovakha distr, 47.51°N, 37.68°E, 25.VIII.2008, KT, 1♂, 2♀♀; Uzbekistan, Karakalpakstan reg., Muynak env. (43.76°N, 59.03°E), VI.1957, V.Sychevskaya, 3♂♂, 9♀♀. Distribution. Southern Palaearctic, from 13°E to 116°E; from 55°N to 35°N.

Lispe manicata Wiedemann, 1830 http://species-id.net/wiki/Lispe_manicata Figs 12, 13, 14

Xenolispa chiragrica Séguy, 1948. Lispe forficata Kurahashi & Shinonaga, 2009: 303 syn. nov. Type locality: Malaysia, Borneo, Sarawak, Bario [3.75°N, 115.45°E].

Material examined. [Indonesia], [Java], Batavia, VI.1908, Jacobson, 1♀ (ZMHU). Cambodia, Sianoukville prov., Ream Nat. Park, 10. 516°N, 103.617°E, 20.IV.2010, O.Kosterin, 1♂. Thailand, Phuket prov., 08.043°N, 98.277°E, 21–26.II.2009, NV, 3♂♂, 2♀♀. Distribution. South-East Asia: Cambodia, Malaysia (Borneo), Indonesia (Java), Singapore (Séguy 1948), S Thailand, S Vietnam (Séguy 1948). 34 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Figure 13–16. Lispe manicata Wiedemann: male (13) male mid tarsi (14) L. glabra Wiedemann: male (15) male mid tarsus (16), df – downcurved fold; vf – “velcro fastener- like setae.

Synonymy. The illustrations of wonderfully modified male mid tarsi and char- acteristic genitalia given by Kurahashi and Shinonaga (2009: fig. 1 c–d, 3 a–d) for L. forficata suggest it’s conspecifity with L. manicata. So, Lispe manicata Wiedemann, 1830 = Lispe forficata Kurahashi & Shinonaga, 2009, syn. n. The characters of female of L. forficata were shortly mentioned by Kurahashi and Shinonaga (2009), but these authors did not compare L. forficata with the most related species L. glabra and, in my opinion, the diagnostically important characters were either not mentioned or given erroneously. Therefore I find it necessary to provide the description of the female below. Description of female. Body length 8.5–9.5 mm. Head. Interfrontalia matt black; frontal triangle brownish-black, subshining, narrow, reaching to lunula. Upper half of fronto-orbital plates brownish-black, subshining, lower part dirty-golden dusted, 3-4 inclinate, 2 reclinate setae and a dense outer row of setulae. Parafacials densely whitish dusted, entirely bare. Gena whitish dusted, about 1.5 times as wide as postpedicel; occiput grey dusted, but less dusted in upper 1/3. Vibrissae strong. Antennae black, long; arista haired on basal half or slightly more, longest aristal hairs half as long as width of antenna. Palpi entirely yellow. Thorax. Pleura and dorsolateral area, including postpronotal lobe and notopleuron, densely grey dusted. Disc of scutum mostly subshining brownish-black, with a pair of thinly dusted vittae situated mesad to dc rows, disc of scutellum subshiny brownish-black. ac hairs in 5-6 irregular rows; dc 0+2, only posterior pair strong (hair-like and indistinct posterior prst dc and 1-2 pairs anterior post dc may be found in some specimens); intraalars 0+1, presutural one absent; supraalars 1+2, the posterior one weak. Katepisternal setae 1+2; anepimeron with 9 (8-10) fine hairs, meron and katepimerom bare. Scutellum bare below at apex. Mesothoracic spiracle yellowish. Wings hyaline, slightly brownish infuscated, vein M distinctly curved forward at apex, calypters whitish, halteres yellow. Revision of the Lispe longicollis-group (Diptera, Muscidae) 35

Legs. Legs long, densely grey dusted; black including coxae, but knees and basal half of t2 and t3 brownish-yellow; f1 with a full row of 6 (5-7) pd setae and with a row of fine pv setulae, but in apical half 2-4 setae in pv row are strong (longer than tibial diameter, stronger than setae in pd row); t1 with p seta, preapical d and apical pd and pv; f2 thickened in basal half, f2 with row of 3-5 a-setae in basal 1/3 and 2 pd at apex; t2 with submedian pd; f3 with short av before middle and shorter than setae of ad row, with a full row of ad subequal to femur depth, preapicals: av and pv; t3 with submedian av, ad and pd setae; hind basitarsus with av seta at base. Abdomen: black with grey pollen. Tergites 1+2 to 5 each with a large blackish spots on dorsal and lateral sides, these spots on tergites 3 and 4 divided at midline by grey dusted interrupted vitta. Male. Similar to female, but mid tarsi modified: apex of mid basitarsus and 4 apical tarsal segments bright yellow; tar2-2 and tar2-3 enlarged, tar2-4 and tar2-5 strongly enlarged; apex of tar2-1 and tar2-2 with long pd-p setae. Cercal plate as shown on Fig. 12.

Lispe microptera Séguy, 1937 http://species-id.net/wiki/Lispe_microptera Figs 3, 8

Material examined. India: Rajasthan state: Jaipur, 26.96°N, 75.85°E, 21–22.II.2011, NV, 10♂♂, 7♀♀; Sambhar salt-lake, 26.916°N, 75.190°E, 23.II.2011, NV, 8♂♂. Distribution. India, Rajasthan and Pakistan, Karachi (type locality). Description of female. Body length 7–7.5 mm, wing length 6mm. Frontal triangle narrow, yellowish dusted; interfrontalia brownish-black. Fronto- orbital plate blackish grey dusted, with 4 inclinate and 2 proclinate setae and dense hairs in outer row. Parafacial covered with hairs. Antenna black, postpedicel short. Arista with hairs two times shorter than antenna width, in apical third bare. Palpus narrow, dirty-yellow. Thorax. Scutum and scutellum brownish dusted with a pair of indistinct vittae, pleura grey dusted. dc 2+4 (medium - medium + medium/weak-medium/weak-strong-strong). Meron with 3-4 setulae above hind coxa, anepimerom with about 15 setulae. Wing with vein R4+5 distinctly curved forward. Legs. Femora dark with yellow apex, tibiae yellow in basal half and dark in apical half, tarsi black. f1 with a complete row of 10-12 pv setae. t1 with submedian p seta. f2 with a row of short a setae in basal half and with 2 pd at apex. t2 with 1 submedian p seta. f3 slightly curved; with a short av seta at basal 1/3 (absent in some specimens) and short pv at apex, av preapical absent. t3 with 1 ad and 1 pd setae at middle. Hind tarsus unmodified. Abdomen grey dusted with large dorsal black spots separated by anteriorly interrupted grey vitta. 36 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Male differs from female as follows: body length 6.5-7 mm, wing length 5-5.5 mm; f2 with a complete row of fine v setae about as long as femur width; f3 in basal half with 4–5 fine long (2–2.5 femur width) pv setae and 1(2) av in basal 1/3; hind tarsus modified: tar3-1 slightly laterally compressed and outward curved, with waved ventral setulae more dense at base and at apex; tar3-2 with waved ventral setulae; male cercal plate as in Fig. 8.

Lispe nuba Wiedemann, 1830 http://species-id.net/wiki/Lispe_nuba

Material examined: Ethiopia: Amhara: Tana Lake env., 1800m asl, 11.54°N, 37.39°E, 2–4.VIII.2012, NV, 6♂♂, 7♀♀; Hayk L., 1920m asl, 11.325°N, 39.688°E, 06.VIII.2012, NV, 3♂♂, 2♀♀; Karakore env., 1500m asl, 10.375°N, 39.933°E, 08.VIII.2012, NV, 4♂♂, 1♀; Oromia: Dedre Zeit, Hora L., 1900m asl, 8.757°N, 38.993°E, 10.VII.2012, NV, 3♂♂, 3♀♀. Egypt: Cairo, 5♂♂, 1♀, with Becker, Kowarz and Hennig determination labels (ZIN). Israel: Yeruham (30.99°N 34.90°E), 22.VII.1962, J.Kugler, 1♂, 1♀ (TAUI). Distribution. Africa and Near East. Remarks. Emden (1941) in the key to African Lispe wrote that in L. nuba “front tibiae without a pv”. It is not correct, all examined specimens have t1 with pv seta in both sexes, though short in males.

Lispe pacifica Shinonaga & Pont, 1992 http://species-id.net/wiki/Lispe_pacifica Figs 17, 18

Material examined. Cambodia, Koh Kong prov., 11.605°N, 103.046°E, XII.2010, NV, 2♀♀. [Taiwan] Formosa: Takao, [22.6N 120.3E], 7.VII.1907, H.Sauter, 6♂♂, 6♀♀, Anping, [23.0N, 120.2E], IX.1908, H.Sauter, 7♂♂, 5♀♀ (ZMHU). Thailand: Chanthaburi prov., Khao Khitchakut env., XI.2009, NV, 1♀; Chi- ang Mai prov., Sop Poeng env., XI.2009, NV, 1♂; Chonburi prov., Pattaya env., XI.2007–XII.2009, NV, 28 ♂♂, ♀♀; Mae Hong Son prov., Pai env., 19.4°N, 98.4°E, 15–25.XI.2010, NV, 19♂♂; Nakhon Ratchasima prov., Khao Yai NP; II.2009; NV, 1♂, 1♀; Phang Nga prov., Khao Lak env., XII.2010, NV, 3♂♂, 1♀; Phuket prov., Nai Thon beach, II.2009; NV, 4♂♂, 1♀; Rayong prov., Ban Phe env., XI.2009, NV, 2♂♂, 2♀♀; Sa Kaew prov., Mueang Sa Kaeo, II.2009; NV, 1♂. Distribution. Widespread in South-East Asia. Revision of the Lispe longicollis-group (Diptera, Muscidae) 37

Ecology

As it was mentioned above, the species of the subgroup 1 of the Lispe longicollis group may be found both on the freshwater and salted basins. L. ethiopica sp.n. was equally common on the freshwater Ziway Lake, on the brackish (2g/l) Langano Lake and on the salt (26g/l) Abijata Lake. L. microptera was collected at brackish lakes and ponds around Jaipur and on the hypersaline (70-300g/l depending on the season) Sambhar Lake. L. longicollis was collected in spring time on the hypersaline Baskunchak Lake, on the seashore salted marshes in Mersin province of Turkey and on the freshwater Titreyen Lake in Antalya province. L. cilitarsis in Israel near Eilat was also found at freshwater of cattle drinking bowl and on a hypersaline lake shore, although in the lat- ter case L. cilitarsis avoided the sites covered with dry salt where only Lispe halophora Becker, 1903 was still present. In contrast to this salt-tolerance, the species of the subgroup 2 of the Lispe longicollis group were observed on freshwater basins only: river banks, rice fields or freshwater lakes/ponds. All species but one prefer open sites, L. manicata seems to be the species of the forest rivers and streams where it was collected in Thailand and Cambodia, the same natural habitat was reported by Kurahashi and Shinonaga (2009) (for L. forficata) in Malaysian Borneo. The species of the Lispe longicollis group mostly feed on slow moving living prey like Nematocera larvae (Fig. 17), but also were observed feeding on dead arthropods (Fig. 18) or even successfully hunting on a small Diptera imago like Paracoenia, Ephy- dridae (Fig. 19) or Syntormon, Dolichopodidae.

Figure 17–19. Feeding. Thailand, Phuket: male L. pacifica: with Chironomidae larva (17) and with dead spider (18) Turkey, Antalya, female L. assimilis with prey - Paracoenia fumosa (Ephydridae) (19).

Correction

I have to apologize for an unfortunate mistake in my previous paper on Lispe taxonomy (Vikhrev 2011, fig. 2): sternite 5 of Lispe draperi Séguy, 1933 was attributed to L. tentaculata (De Geer, 1776) and vice versa. 38 Nikita E. Vikhrev / ZooKeys 235: 23–39 (2012)

Acknowledgements

I thank Adrian Pont (Oxford) for his various help and for the BMNH material, Oleg Kosterin (Novosibirsk) and Dmitry Gavryushin (Moscow) for their corrections of the text, Amnon Freidberg (Tel-Aviv), Joachim Ziegler (Berlin) and Emilia Narchuk (St Petersburg) for the very interesting material from TAUI, ZMHU and ZIN respectively.

References

Emden FI van (1941) Keys to the Muscidae of the Ethiopian region: Scatophaginae, An- thomyiinae, Lispinae, Fanniinae. Bulletin of Entomological Research 32: 251–275. doi: 10.1017/S0007485300017193 Hennig W (1960) Family Muscidae (Lieferung 209 and 213). In: Lindner E (Ed.) Die Fliegen der Palaarktischen Region. Stuttgart 63b, 385–480. Kurahashi H, Shinonaga S (2009) Two new species of the Muscid flies from Sarawak, East Ma- laysia (Diptera, Muscidae), Japanese Journal of Systematic Entomology 15(2): 299–306. Pont AC (1977) Family Muscidae. In: Delfinado MD, Hardy DE (Eds) Catalogue of the Dip- tera of the Oriental Region 3. University Press of Hawaii, Honolulu, 451–523. Pont AC (1980) Family Muscidae. In: Crosskey RW (Ed.) Catalogue of the Diptera of the Afrotropical Region. British Museum (Natural History), London, 721–761. Pont AC (1986) Family Muscidae. In: Soós A, Papp L (Eds) Catalogue of Palaearctic Diptera 11. Akadémia Kiadó, Budapest, 57–215. Pont AC (1991) A review of the Fanniidae and Muscidae of the Arabian Peninsula. Fauna of Saudi Arabia 12: 312–365. Pont AC, Werner D (2006) The Types of Fanniidae and Muscidae (Diptera) in the Mu- seum für Naturkunde, Humboldt-Universität zu Berlin, Germany. Mitteilungen aus dem Museum für Naturkunde in Berlin – Zoologische Reihe 82(1): 3–139. doi: 10.1002/ mmnz.200600001 Séguy E (1948) Dipteres nouveaux ou peu connus d’extreme-Orient. Notes d’Entomologie Chinoise 12(14): 153–172. Shinonaga S, Pont AC (1992) A new species of the genus Lispe Latreille, with notes on two related species, L. assimilis Wiedemann and L. microptera Séguy (Diptera, Muscidae). Japa- nese journal of entomology 60(4): 715–722. Stein P (1900) Einige dem Genueser Museum gehorige aus Neu-Guinea und Umgegend stam- mende Anthomyiden. Annali del Museo Civico di Storia Naturale di Genova 40: 374–95. Stein P (1908) Zoologische und anthropologische Ergebnisse einer Forschungsreise im westli- chen und zentralen Sudafrika ausgefuhrt in den Jahren 1903-1905 mit Unterstutzung der Kgl. Preussischen Akademie der Wissenschaften zu Berlin von Dr. Leonhard Schultze, Pro- fessor der Erdkunde an der Universitat Jena. Erster Band: Systematik und Tiergeographie. IV. Insecta (Erste Serie). D. Diptera. 4. Anthomyidae. – Denkschriften der medizinisch- naturwissenschaftlichen Gesellschaft zu Jena 13: 171–174. Stein P (1913) Neue afrikanische Anthomyiden. Annales historico-naturales Musei nationalis Hungarici 11: 457–583 Stein P (1918) Zur weitern Kenntnis aussereuropaeischer Anthomyiden. Annales Historico- Naturales Musei Nationalis Hungarici 16: 147–244. Revision of the Lispe longicollis-group (Diptera, Muscidae) 39

Vikhrev N (2011) Review of the Palaearctic members of the Lispe tentaculata species-group (Diptera, Muscidae): revised key, synonymy and notes on ecology. ZooKeys 84: 59–70. doi: 10.3897/zookeys.84.819